It has come to my attention

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It has come to my attention

Postby Eddie » Sun Dec 20, 2009 2:50 am

That there was a murder clue in a dead man's eye.
This also make no sense, because;
Death
From Wikipedia, the free encyclopedia
Jump to:navigation, search
For other uses of "death" and "deceased", see Death (disambiguation).
Universal symbol for death: a human skull
A dead Confederate soldier sprawled out in Petersburg, Virginia, 1865, during the American Civil War

Death is the termination of the biological functions that define a living organism. It refers both to a particular event and to the condition that results thereby. The true nature of the latter has for millennia been a central concern of the world's religious traditions and of philosophical enquiry. Belief in some kind of afterlife or rebirth is a central aspect of many religious traditions. Within the scientific community, many suppose death to terminate mind or consciousness. The effect of physical death on any possible mind or soul remains for many an open question. Cognitive science has yet to explain fully the origin and nature of consciousness; any view about the existence or non-existence of consciousness after death remains speculative.[1][2]

Humans and the vast majority of other animals die in due course from senescence. Remarkable exceptions include the hydra, and the jellyfish turritopsis nutricula, which is thought to possess in effect biological immortality.[3]

Intervening phenomena which commonly bring about death earlier include malnutrition, disease, or accidents resulting in terminal physical injury. Predation is a cause of death for many species. Intentional human activity causing death includes suicide, homicide, and war. Roughly 150,000 people die each day across the globe.[4] Death in the natural world can also occur as an indirect result of human activity: an increasing cause of species depletion in recent times has been destruction of ecosystems as a consequence of the widening spread of industrial technology.[5]

Physiological death is now seen as less an event than a process: conditions once considered indicative of death are now reversible.[6] Where in the process a dividing line is drawn between life and death depends on factors beyond the presence or absence of vital signs. In general, clinical death is neither necessary nor sufficient for a determination of legal death. A patient with working heart and lungs determined to be brain dead can be pronounced legally dead without clinical death occurring. Precise medical definition of death, in other words, becomes more problematic, paradoxically, as scientific knowledge and technology advance.
Contents
[hide]

* 1 Signs and symptoms
* 2 Diagnosis
o 2.1 Problems of definition
o 2.2 Legal
o 2.3 Misdiagnosed
* 3 Causes
o 3.1 Autopsy
* 4 Prevention
* 5 Society and culture
* 6 In biology
o 6.1 Natural selection
o 6.2 Extinction
o 6.3 Evolution of aging
* 7 See also
* 8 References
* 9 Further reading
* 10 External links

[edit] Signs and symptoms

Signs of death, or strong indications that a person is no longer alive are:

* Ceasing respiration
* The body no longer metabolises
* Pallor mortis, paleness which happens in the 15–120 minutes after the death
* Livor mortis, a settling of the blood in the lower (dependent) portion of the body
* Algor mortis, the reduction in body temperature following death. This is generally a steady decline until matching ambient temperature
* Rigor mortis, the limbs of the corpse become stiff (Latin rigor) and difficult to move or manipulate
* Decomposition, the reduction into simpler forms of matter

[edit] Diagnosis
[edit] Problems of definition
What is death? A flower, a skull and an hour-glass stand in for Life, Death and Time in this 17th Century painting by Philippe de Champaigne

For those who define death as a state following the state of life, one of the challenges in defining death is in distinguishing it from life. Death would seem to refer to either the moment at which life ends, or when the state that follows life begins. However, determining when death has occurred requires drawing precise conceptual boundaries between life and death. This is problematic however because there is little consensus over how to define life. Some have suggested defining life in terms of consciousness. When consciousness ceases, a living organism can be said to have died. One of the notable flaws in this approach is that there are many organisms which are alive but probably not conscious (for example, single-celled organisms). Another problem with this approach is in defining consciousness, which remains a mystery to modern scientists, psychologists and philosophers. This general problem of defining death applies to the particular challenge of defining death in the context of medicine.

Other definitions for death focus on the character of cessation of something.[7] In this context 'death' describes merely the state where something has ceased, e.g., life. Thus, the definition of 'life' simultaneously defines death.

Historically, attempts to define the exact moment of a human's death have been problematic. Death was once defined as the cessation of heartbeat (cardiac arrest) and of breathing, but the development of CPR and prompt defibrillation have rendered that definition inadequate because breathing and heartbeat can sometimes be restarted. Events which were causally linked to death in the past no longer kill in all circumstances; without a functioning heart or lungs, life can sometimes be sustained with a combination of life support devices, organ transplants and artificial pacemakers.

Today, where a definition of the moment of death is required, doctors and coroners usually turn to "brain death" or "biological death" to define a person as being clinically dead; people are considered dead when the electrical activity in their brain ceases. It is presumed that an end of electrical activity indicates the end of consciousness. However, suspension of consciousness must be permanent, and not transient, as occurs during certain sleep stages, and especially a coma. In the case of sleep, EEGs can easily tell the difference.

However, the category of "brain death" is seen by some scholars to be problematic. For instance, Dr Franklin Miller, senior faculty member at the Department of Bioethics, National Institutes of Health, notes "By the late 1990s, however, the equation of brain death with death of the human being was increasingly challenged by scholars, based on evidence regarding the array of biological functioning displayed by patients correctly diagnosed as having this condition who were maintained on mechanical ventilation for substantial periods of time. These patients maintained the ability to sustain circulation and respiration, control temperature, excrete wastes, heal wounds, fight infections and, most dramatically, to gestate fetuses (in the case of pregnant "brain-dead" women)." [8]

Many have challenged the idea that brain death is equivalent to the cessation of consciousness. Critics point out that much of human consciousness is embodied in numerous body parts and that the end of electrical impulses in the brain does not necessarily indicate that this embodied consciousness has also ceased. Given this possibility, brain death does not necessitate the end of consciousness, and thus brain dead people may still be alive. Furthermore, some have argued, even if brain death does mean the end of consciousness for a human being, the whole notion that cessation of consciousness indicates death is problematic. Critics note the existence of many single-celled organisms such as bacteria that we consider to be alive but which many doubt are conscious. If life does not require consciousness, defining death in terms of "brain death" is a dubious procedure, even if the brain is the seat of consciousness. Thus while legal concerns surrounding death force us to develop a working definition of death, it is not at all clear that the current American definition, according to brain death, coincides at all with a definition that can be reasonably endorsed.[who?]

Those people maintaining that only the neo-cortex of the brain is necessary for consciousness sometimes argue that only electrical activity there should be considered when defining death. Eventually it is possible that the criterion for death will be the permanent and irreversible loss of cognitive function, as evidenced by the death of the cerebral cortex. All hope of recovering human thought and personality is then gone given current and foreseeable medical technology. However, at present, in most places the more conservative definition of death — irreversible cessation of electrical activity in the whole brain, as opposed to just in the neo-cortex — has been adopted (for example the Uniform Determination Of Death Act in the United States). In 2005, the Terri Schiavo case brought the question of brain death and artificial sustenance to the front of American politics.

Even by whole-brain criteria, the determination of brain death can be complicated. EEGs can detect spurious electrical impulses, while certain drugs, hypoglycemia, hypoxia, or hypothermia can suppress or even stop brain activity on a temporary basis. Because of this, hospitals have protocols for determining brain death involving EEGs at widely separated intervals under defined conditions.
[edit] Legal
See also: Legal death

In the United States, a person is dead by law if a Statement of Death or Death Certificate is approved by a licensed medical practitioner. Various legal consequences follow death, including the removal from the person of what in legal terminology is called personhood.

The possession of brain activities, or ability to resume brain activity, is a necessary condition to legal personhood in the United States. "It appears that once brain death has been determined … no criminal or civil liability will result from disconnecting the life-support devices." (Dority v. Superior Court of San Bernardino County, 193 Cal.Rptr. 288, 291 (1983))
[edit] Misdiagnosed

There are many anecdotal references to people being declared dead by physicians and then 'coming back to life', sometimes days later in their own coffin, or when embalming procedures are just about to begin. Owing to significant scientific advancements in the Victorian era, some people in Britain became obsessively worried about living after being declared dead.[9]

In cases of electric shock, CPR for an hour or longer can allow stunned nerves to recover, allowing an apparently dead person to survive. People found unconscious under icy water may survive if their faces are kept continuously cold until they arrive at an emergency room.[10] This "diving response", in which metabolic activity and oxygen requirements are minimal, is something humans share with cetaceans called the mammalian diving reflex.[10]

As medical technologies advance, ideas about when death occurs may have to be re-evaluated in light of the ability to restore a person to vitality after longer periods of apparent death (as happened when CPR and defibrillation showed that cessation of heartbeat is inadequate as a decisive indicator of death). The lack of electrical brain activity may not be enough to consider someone scientifically dead. Therefore, the concept of information theoretical death has been suggested as a better means of defining when true death actually occurs, though the concept has few practical applications outside of the field of cryonics.

There have been some scientific attempts to bring dead organisms back to life, but with limited success.[11] In science fiction scenarios where such technology is readily available, real death is distinguished from reversible death.
[edit] Causes
See also: List of causes of death by rate and List of preventable causes of death
The body of Pope John Paul II lying in state in St. Peter's Basilica, 2005

The leading cause of death in developing countries is infectious disease. The leading causes of death in developed countries are atherosclerosis (heart disease and stroke), cancer, and other diseases related to obesity and aging. These conditions cause loss of homeostasis, leading to cardiac arrest, causing loss of oxygen and nutrient supply, causing irreversible deterioration of the brain and other tissues. Of the roughly 150,000 people who die each day across the globe, about two thirds — 100,000 per day — die of age-related causes.[4] In industrialized nations, the proportion is much higher, reaching 90%.[4] With improved medical capability, dying has become a condition to be managed. Home deaths, once normal, are now rare in the developed world.

In developing nations, inferior sanitary conditions and lack of access to modern medical technology makes death from infectious diseases more common than in developed countries. One such disease is tuberculosis, a bacterial disease which killed 1.7 million people in 2004.[12] Malaria causes about 400–900 million cases of fever and approximately one to three million deaths annually.[13] AIDS death toll in Africa may reach 90-100 million by 2025.[14][15]

According to Jean Ziegler, who was the United Nations Special reporter on the Right to Food from 2000 to March 2008; mortality due to malnutrition accounted for 58% of the total mortality rate in 2006. Ziegler says worldwide approximately 62 million people died from all causes and of those deaths more than 36 million died of hunger or diseases due to deficiencies in micronutrients."[16]

Tobacco smoking killed 100 million people worldwide in the 20th century and could kill 1 billion people around the world in the 21st century, a WHO Report warned.[17][18]

Many leading developed world causes of death can be postponed by diet and physical activity, but the accelerating incidence of disease with age still imposes limits on human longevity. The evolutionary cause of aging is, at best, only just beginning to be understood. It has been suggested that direct intervention in the aging process may now be the most effective intervention against major causes of death.[19]
[edit] Autopsy
Rembrandt turns an autopsy into a masterpiece: The Anatomy Lesson of Dr. Nicolaes Tulp

An autopsy, also known as a postmortem examination or an obduction, is a medical procedure that consists of a thorough examination of a human corpse to determine the cause and manner of a person's death and to evaluate any disease or injury that may be present. It is usually performed by a specialized medical doctor called a pathologist.

Autopsies are either performed for legal or medical purposes. A forensic autopsy is carried out when the cause of death may be a criminal matter, while a clinical or academic autopsy is performed to find the medical cause of death and is used in cases of unknown or uncertain death, or for research purposes. Autopsies can be further classified into cases where external examination suffices, and those where the body is dissected and an internal examination is conducted. Permission from next of kin may be required for internal autopsy in some cases. Once an internal autopsy is complete the body is generally reconstituted by sewing it back together. Autopsy is important in a medical environment and may shed light on mistakes and help improve practices.

A "necropsy" is an older term for a postmortem examination, unregulated, and not always a medical procedure. In modern times the term is more often used in the postmortem examination of the corpses of animals.
[edit] Prevention
Main article: Life extension

Life extension refers to an increase in maximum or average lifespan, especially in humans, by slowing down or reversing the processes of aging. Average lifespan is determined by vulnerability to accidents and age or lifestyle-related afflictions such as cancer, or cardiovascular disease. Extension of average lifespan can be achieved by good diet, exercise and avoidance of hazards such as smoking. Maximum lifespan is determined by the rate of aging for a species inherent in its genes. Currently, the only widely recognized method of extending maximum lifespan is calorie restriction. Theoretically, extension of maximum lifespan can be achieved by reducing the rate of aging damage, by periodic replacement of damaged tissues, or by molecular repair or rejuvenation of deteriorated cells and tissues.

Researchers of life extension are a subclass of biogerontologists known as "biomedical gerontologists". They try to understand the nature of aging and they develop treatments to reverse aging processes or to at least slow them down, for the improvement of health and the maintenance of youthful vigor at every stage of life. Those who take advantage of life extension findings and seek to apply them upon themselves are called "life extensionists" or "longevists". The primary life extension strategy currently is to apply available anti-aging methods in the hope of living long enough to benefit from a complete cure to aging once it is developed, which given the rapidly advancing state of biogenetic and general medical technology, could conceivably occur within the lifetimes of people living today.
[edit] Society and culture
Main article: Death and culture
Death haunts even the beautiful: an early 20th century artist says, "All is Vanity"

Death is the center of many traditions and organizations, and is a feature of every culture around the world. Much of this revolves around the care of the dead, as well as the afterlife and the disposal of bodies upon the onset of death. The disposal of human corpses does, in general, begin with the last offices before significant time has passed, and ritualistic ceremonies often occur, most commonly interment or cremation. This is not a unified practice, however, as in Tibet for instance the body is given a sky burial and left on a mountain top. Proper preparation for death and techniques and ceremonies for producing the ability to transfer one's spiritual attainments into another body (reincarnation) are subjects of detailed study in Tibet.[20] Mummification or embalming is also prevalent in some cultures, to retard the rate of decay.

Legal aspects of death are also part of many cultures, particularly the settlement of the deceased estate and the issues of inheritance and in some countries, inheritance taxation.

Capital punishment is also a divisive aspect of death in culture. In most places that practice capital punishment today, the death penalty is reserved as punishment for premeditated murder, espionage, treason, or as part of military justice. In some countries, sexual crimes, such as adultery and sodomy, carry the death penalty, as do religious crimes such as apostasy, the formal renunciation of one's religion. In many retentionist countries, drug trafficking is also a capital offense. In China human trafficking and serious cases of corruption are also punished by the death penalty. In militaries around the world courts-martial have imposed death sentences for offenses such as cowardice, desertion, insubordination, and mutiny.[21]

Death in warfare and in suicide attack also have cultural links, and the ideas of dulce et decorum est pro patria mori, mutiny punishable by death, grieving relatives of dead soldiers and death notification are embedded in many cultures. Recently in the western world, with the supposed increase in terrorism following the September 11 attacks, but also further back in time with suicide bombings, kamikaze missions in World War II and suicide missions in a host of other conflicts in history, death for a cause by way of suicide attack, and martyrdom have had significant cultural impacts.

Suicide in general, and particularly euthanasia are also points of cultural debate. Both acts are understood very differently in contrasting cultures. In Japan, for example, ending a life with honor by seppuku was considered a desirable death, whereas according to traditional Christian and Islamic cultures, suicide is viewed as a sin. Death is personified in many cultures, with such symbolic representations as the Grim Reaper, Azrael and Father Time.
[edit] In biology
[edit] Natural selection
Main articles: competition (biology), natural selection, and extinction

Contemporary evolutionary theory sees death as an important part of the process of natural selection. It is considered that organisms less adapted to their environment are more likely to die having produced fewer offspring, thereby reducing their contribution to the gene pool. Their genes are thus eventually bred out of a population, leading at worst to extinction and, more positively, making possible the process referred to as speciation. Frequency of reproduction plays an equally important role in determining species survival: an organism that dies young but leaves numerous offspring displays, according to Darwinian criteria, much greater fitness than a long-lived organism leaving only one.
[edit] Extinction
Main article: Extinction
Dead as a dodo: the bird that became a byword in English for species extinction [22]

Extinction is the cessation of existence of a species or group of taxa, reducing biodiversity. The moment of extinction is generally considered to be the death of the last individual of that species (although the capacity to breed and recover may have been lost before this point). Because a species' potential range may be very large, determining this moment is difficult, and is usually done retrospectively. This difficulty leads to phenomena such as Lazarus taxa, where a species presumed extinct abruptly "reappears" (typically in the fossil record) after a period of apparent absence. New species arise through the process of speciation, an aspect of evolution. New varieties of organisms arise and thrive when they are able to find and exploit an ecological niche — and species become extinct when they are no longer able to survive in changing conditions or against superior competition. After death the remains of an organism become part of the biogeochemical cycle. Animals may be consumed by a predator or a scavenger. Organic material may then be further decomposed by detritivores, organisms which recycle detritus, returning it to the environment for reuse in the food chain. Examples of detritivores include earthworms, woodlice and dung beetles.

Microorganisms also play a vital role, raising the temperature of the decomposing matter as they break it down into yet simpler molecules. Not all materials need be decomposed fully, however. Coal, a fossil fuel formed over vast tracts of time in swamp ecosystems, is one example.
[edit] Evolution of aging
Main article: Evolution of ageing

Inquiry into the evolution of aging aims to explain why so many living things and the vast majority of animals weaken and die with age (a notable exception being hydra, which may be biologically immortal). The evolutionary origin of senescence remains one of the fundamental puzzles of biology. Gerontology specializes in the science of human aging processes.
[edit] See also

* Afterlife
* Bardo Thodol (Tibetan Book of the Dead)
* Burial
* Black Death
* Cadaveric spasm
* Death, Desire and Loss in Western Culture by Jonathan Dollimore
* Dead bell
* Death drive
* Death erection
* Death messengers
* Death (personification)
* Death rattle
* Día de los Muertos (Day of the Dead)
* Dying declaration
* International Necronautical Society



* Karōshi
* Last rites
* Leading preventable causes of death
* Legal death
* List of causes of death by rate
* List of natural disasters by death toll
* List of wars and disasters by death toll
* Mortician
* Near-death experience
* Post Mortem Interval
* Pseudocide
* Thanatology
* Vampire
* World War I casualties
* World War II casualties
* Zombie

[edit] References

1. ^ "The Big Questions: What is consciousness?". New Scientist. 18 November 2006. http://www.newscientist.com/article/mg1 ... ?full=true. Retrieved 2009-09-04.
2. ^ "Facing up to the problem of consciousness". http://consc.net/papers/facing.html.
3. ^ Guerin, John C. (2009). Emerging Area of Aging Research: Long-lived Animals with "Negligible Senescence". http://www.agelessanimals.org/. Retrieved August 21, 2009.
4. ^ a b c Aubrey D.N.J, de Grey (2007). "Life Span Extension Research and Public Debate: Societal Considerations" (PDF). Studies in Ethics, Law, and Technology 1 (1, Article 5). doi:10.2202/1941-6008.1011. http://www.mfoundation.org/files/sens/ENHANCE-PP.pdf. Retrieved March 20, 2009.
5. ^ Human Activities Cause of Current Extinction Crisis, accessed 7 April 2009
6. ^ Crippen, David. "Brain Failure and Brain Death". ACS Surgery Online, Critical Care, April 2005. Archived from the original on 24 June 2006. http://web.archive.org/web/200606241324 ... cs0812.htm. Retrieved 2007-01-09.
7. ^ Oxford English Dictionary
8. ^ FG Miller "Death and organ donation: back to the future" Journal of Medical Ethics 2009;35:616-620
9. ^ As reflected from at least one article of literature by authors like Edgar Allan Poe, where subjects were buried alive.
10. ^ a b Limmer, D. et al. (2006). Emergency care (AHA update, Ed. 10e). Prentice Hall.
11. ^ Blood Swapping Reanimates Dead Dogs
12. ^ World Health Organization (WHO). Tuberculosis Fact sheet N°104 - Global and regional incidence. March 2006, Retrieved on 6 October 2006.
13. ^ USAID’s Malaria Programs
14. ^ Aids could kill 90 million Africans, says UN
15. ^ AIDS Toll May Reach 100 Million in Africa, Washington Post
16. ^ Jean Ziegler, L'Empire de la honte, Fayard, 2007 ISBN 978-2-253-12115-2 p.130.
17. ^ Tobacco Could Kill One Billion By 2100, World Health Organization Report Warns
18. ^ Tobacco could kill more than 1 billion this century: World Health Organization
19. ^ SJ Olshanksy et al. (2006). "Longevity dividend: What should we be doing to prepare for the unprecedented aging of humanity?". The Scientist 20: 28–36. http://www.grg.org/resources/TheScientist.htm. Retrieved 2007-03-31.
20. ^ Mullin (1999).
21. ^ "Shot at Dawn, campaign for pardons for British and Commonwealth soldiers executed in World War I". Shot at Dawn Pardons Campaign. http://www.shotatdawn.org.uk/. Retrieved 2006-07-20.
22. ^ Diamond, Jared (1999). "Up to the Starting Line". Guns, Germs, and Steel. W. W. Norton. pp. 43–44. ISBN 0-393-31755-2.

[edit] Further reading

* Appel, JM. Defining Death: When Physicians and Families Differ. Journal of Medical Ethics Fall 2005.
* Child AM (1995) J Archaeolog Sci 22: 165-174it funny
* Mullin, Glenn H. (1998). Living in the Face of Death: The Tibetan Tradition. 2008 reprint: Snow Lion Publications, Ithica, New York. ISBN 978-1-55939-310-2.
* Piepenbrink H (1985) J Archaeolog Sci 13: 417-430
* Piepenbrink H (1989) Applied Geochem 4: 273-280
* Pounder, Derrick J. (2005-12-15). "Postmortem changes and time of death". University of Dundee. http://www.dundee.ac.uk/forensicmedicin ... edeath.pdf. Retrieved 2006-12-13.
* Vass AA (2001) Microbiology Today 28: 190-192 at: [1]

[edit] External links
Search Wikiquote Wikiquote has a collection of quotations related to: Death
Search Wikimedia Commons Wikimedia Commons has media related to: Death
Search Wiktionary Look up death in Wiktionary, the free dictionary.

* Death at the Open Directory Project
* Death (Stanford Encyclopedia of Philosophy)
* Doctors Change the Way They Think About Death
* Odds of dying from various injuries or accidents Source: National Safety Council, United States, 2001
* Causes of Death
* Causes of Death 1916 How the medical profession categorized causes of death a century ago.
* George Wald: The Origin of Death A biologist explains life and death in different kinds of organisms in relation to evolution.
* Before and After Death Interviews with people dying in hospices, and portraits of them before, and shortly after, death


Preceded by
Old age Stages of human development
Death Succeeded by
Decomposition

Eye
From Wikipedia, the free encyclopedia
Jump to:navigation, search
For information specific to humans, see Human eye.
For other uses, see Eye (disambiguation).
Eye
Schematic diagram of the human eye en.svg
Schematic diagram of the vertebrate eye.
Krilleyekils.jpg
Compound eye of Antarctic krill

Eyes are organs that detect light, and send electrical impulses along the optic nerve to the visual and other areas of the brain. Complex optical systems with resolving power have come in ten fundamentally different forms, and 96% of animal species possess a complex optical system.[1] Image-resolving eyes are present in cnidaria, molluscs, chordates, annelids and arthropods.[2]

The simplest "eyes", such as those in unicellular organisms, do nothing but detect whether the surroundings are light or dark, which is sufficient for the entrainment of circadian rhythms. From more complex eyes, retinal photosensitive ganglion cells send signals along the retinohypothalamic tract to the suprachiasmatic nuclei to effect circadian adjustment.
Contents
[hide]

* 1 Overview
* 2 Evolution
* 3 Types of eye
o 3.1 Simple eyes
+ 3.1.1 Pit eyes
+ 3.1.2 Pinhole eye
+ 3.1.3 Spherical lensed eye
# 3.1.3.1 Weaknesses
+ 3.1.4 Multiple lenses
+ 3.1.5 Refractive cornea
+ 3.1.6 Reflector eyes
o 3.2 Compound eyes
+ 3.2.1 Apposition eyes
+ 3.2.2 Superposition eyes
+ 3.2.3 Parabolic superposition
+ 3.2.4 Other
* 4 Relationship to lifestyle
* 5 Acuity
* 6 Color
* 7 Rods and cones
* 8 Pigment
* 9 See also
* 10 References
* 11 External links

[edit] Overview
Eye of the wisent,
the European bison

Complex eyes can distinguish shapes and colors. The visual fields of many organisms, especially predators, involve large areas of binocular vision to improve depth perception; in other organisms, eyes are located so as to maximise the field of view, such as in rabbits and horses, which have monocular vision.

The first proto-eyes evolved among animals 540 million years ago, about the time of the Cambrian explosion.[citation needed] The last common ancestor of animals possessed the biochemical toolkit necessary for vision, and more advanced eyes have evolved in 96% of animal species in 6 of the thirty-plus[note 1] main phyla.[1] In most vertebrates and some molluscs, the eye works by allowing light to enter it and project onto a light-sensitive panel of cells, known as the retina, at the rear of the eye. The cone cells (for color) and the rod cells (for low-light contrasts) in the retina detect and convert light into neural signals for vision. The visual signals are then transmitted to the brain via the optic nerve. Such eyes are typically roughly spherical, filled with a transparent gel-like substance called the vitreous humour, with a focusing lens and often an iris; the relaxing or tightening of the muscles around the iris change the size of the pupil, thereby regulating the amount of light that enters the eye,[3] and reducing aberrations when there is enough light.[4]

The eyes of cephalopods, fish, amphibians and snakes usually have fixed lens shapes, and focusing vision is achieved by telescoping the lens — similar to how a camera focuses.[5]

Compound eyes are found among the arthropods and are composed of many simple facets which, depending on the details of anatomy, may give either a single pixelated image or multiple images, per eye. Each sensor has its own lens and photosensitive cell(s). Some eyes have up to 28,000 such sensors, which are arranged hexagonally, and which can give a full 360-degree field of vision. Compound eyes are very sensitive to motion. Some arthropods, including many Strepsiptera, have compound eyes of only a few facets, each with a retina capable of creating an image, creating vision. With each eye viewing a different thing, a fused image from all the eyes is produced in the brain, providing very different, high-resolution images.

Possessing detailed hyperspectral color vision, the Mantis shrimp has been reported to have the world's most complex color vision system.[6] Trilobites, which are now extinct, had unique compound eyes. They used clear calcite crystals to form the lenses of their eyes. In this, they differ from most other arthropods, which have soft eyes. The number of lenses in such an eye varied, however: some trilobites had only one, and some had thousands of lenses in one eye.

In contrast to compound eyes, simple eyes are those that have a single lens. For example, jumping spiders have a large pair of simple eyes with a narrow field of view, supported by an array of other, smaller eyes for peripheral vision. Some insect larvae, like caterpillars, have a different type of simple eye (stemmata) which gives a rough image. Some of the simplest eyes, called ocelli, can be found in animals like some of the snails, which cannot actually "see" in the normal sense. They do have photosensitive cells, but no lens and no other means of projecting an image onto these cells. They can distinguish between light and dark, but no more. This enables snails to keep out of direct sunlight. In organisms dwelling near deep-sea vents, compound eyes have been secondarily simplified and adapted to spot the infra-red light produced by the hot vents - in this way the bearers can spot hot springs and avoid being boiled alive.[7]
[edit] Evolution
Main article: Evolution of the eye
Diagram of eye evolution.svg

The common origin (monophyly) of all animal eyes is now widely accepted as fact based on shared anatomical and genetic features of all eyes; that is, all modern eyes, varied as they are, have their origins in a proto-eye believed to have evolved some 540 million years ago.[8][9][10] The majority of the advancements in early eyes are believed to have taken only a few million years to develop, as the first predator to gain true imaging would have touched off an "arms race".[11] Prey animals and competing predators alike would be at a distinct disadvantage without such capabilities and would be less likely to survive and reproduce. Hence multiple eye types and subtypes developed in parallel.

Eyes in various animals show adaption to their requirements. For example, birds of prey have much greater visual acuity than humans, and some can see ultraviolet light. The different forms of eye in, for example, vertebrates and mollusks are often cited as examples of parallel evolution, despite their distant common ancestry.

The earliest eyes, called "eyespots", were simple patches of photoreceptor cells, physically similar to the receptor patches for taste and smell. These eyespots could only sense ambient brightness: they could distinguish light and dark, but not the direction of the lightsource.[12] This gradually changed as the eyespot depressed into a shallow "cup" shape, granting the ability to slightly discriminate directional brightness by using the angle at which the light hit certain cells to identify the source. The pit deepened over time, the opening diminished in size, and the number of photoreceptor cells increased, forming an effective pinhole camera that was capable of slightly distinguishing dim shapes.[13]

The thin overgrowth of transparent cells over the eye's aperture, originally formed to prevent damage to the eyespot, allowed the segregated contents of the eye chamber to specialize into a transparent humour that optimized colour filtering, blocked harmful radiation, improved the eye's refractive index, and allowed functionality outside of water. The transparent protective cells eventually split into two layers, with circulatory fluid in between that allowed wider viewing angles and greater imaging resolution, and the thickness of the transparent layer gradually increased, in most species with the transparent crystallin protein.[14]

The gap between tissue layers naturally formed a bioconvex shape, an optimally ideal structure for a normal refractive index. Independently, a transparent layer and a nontransparent layer split forward from the lens: the cornea and iris. Separation of the forward layer again forms a humour, the aqueous humour. This increases refractive power and again eases circulatory problems. Formation of a nontransparent ring allows more blood vessels, more circulation, and larger eye sizes.[14]
[edit] Types of eye

Nature has produced ten different eye layouts — indeed every way of capturing an image has evolved at least once in nature, with the exceptions of zoom and Fresnel lenses. Eye types can be categorized into "simple eyes", with one concave chamber, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface.[1] Note that "simple" does not imply a reduced level of complexity or acuity. Indeed, any eye type can be adapted for almost any behaviour or environment. The only limitations specific to eye types are that of resolution — the physics of compound eyes prevents them from achieving a resolution better than 1°. Also, superposition eyes can achieve greater sensitivity than apposition eyes, so are better suited to dark-dwelling creatures.[1] Eyes also fall into two groups on the basis of their photoreceptor's cellular construction, with the photoreceptor cells either being cilliated (as in the vertebrates) or rhabdomic. These two groups are not monophyletic; the cnidaira also possess cilliated cells, [15] and some annelids possess both.[16]
[edit] Simple eyes

Simple eyes are rather ubiquitous, and lens-bearing eyes have evolved at least seven times; in vertebrates, cephalopods, annelids, crustacea and cubozoa.[17]
[edit] Pit eyes

Pit eyes, also known as stemma, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eyespot, to allow the organism to deduce the angle of incoming light.[1] Found in about 85% of phyla, these basic forms were probably the precursors to more advanced types of "simple eye". They are small, comprising up to about 100 cells covering about 100 µm.[1] The directionality can be improved by reducing the size of the aperture, by incorporating a reflective layer behind the receptor cells, or by filling the pit with a refractile material.[1]
[edit] Pinhole eye
Nautiluses bear a pinhole eye

The pinhole eye is an "advanced" form of pit eye incorporating several improvements, most notably a small aperture (which may be adjustable) and deep pit. It is only found in the nautiloids.[1] Without a lens to focus the image, it produces a blurry image, and will blur out a point to the size of the aperture. Consequently, nautiloids can't discriminate between objects with an angular separation of less than 11°.[1] Shrinking the aperture would produce a sharper image, but let in less light.[1]
[edit] Spherical lensed eye

The resolution of pit eyes can be greatly improved by incorporating a material with a higher refractive index to form a lens, which may greatly reduce the blur radius encountered — hence increasing the resolution obtainable.[1] The most basic form, still seen in some gastropods and annelids, consists of a lens of one refractive index. A far sharper image can be obtained using materials with a high refractive index, decreasing to the edges — this decreases the focal length and thus allows a sharp image to form on the retina.[1] This also allows a larger aperture for a given sharpness of image, allowing more light to enter the lens; and a flatter lens, reducing spherical aberration.[1] Such an inhomogeneous lens is necessary in order for the focal length to drop from about 4 times the lens radius, to 2.5 radii.[1]

Heterogeneous eyes have evolved at least eight times — four or more times in gastropods, once in the copepods, once in the annelids and once in the cephalopods.[1] No aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown".[1]

This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimize the effect of eye motion while the animal moves, most such eyes have stabilizing eye muscles.[1]

The ocelli of insects bear a simple lens, but their focal point always lies behind the retina; consequently they can never form a sharp image. This capitulates the function of the eye. Ocelli (pit-type eyes of arthropods) blur the image across the whole retina, and are consequently excellent at responding to rapid changes in light intensity across the whole visual field — this fast response is further accelerated by the large nerve bundles which rush the information to the brain.[18] Focusing the image would also cause the sun's image to be focused on a few receptors, with the possibility of damage under the intense light; shielding the receptors would block out some light and thus reduce their sensitivity.[18] This fast response has led to suggestions that the ocelli of insects are used mainly in flight, because they can be used to detect sudden changes in which way is up (because light, especially UV light which is absorbed by vegetation, usually comes from above).[18]
[edit] Weaknesses

One weakness of this eye construction is that chromatic aberration is still quite high[1] — although for organisms without color vision, this is a very minor concern.

A weakness of the vertebrate eye is the blind spot at the optic disc where the optic nerve is formed at the back of the eye; there are no light sensitive rods or cones to respond to a light stimulus at this point. By contrast, the cephalopod eye has no blind spot as the retina is in the opposite orientation.
[edit] Multiple lenses

Some marine organisms bear more than one lens; for instance the copepod Pontella has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another copepod, Copilia's eyes have two lenses, arranged like those in a telescope.[1] Such arrangements are rare and poorly understood, but represent an interesting alternative construction. An interesting use of multiple lenses is seen in some hunters such as eagles and jumping spiders, which have a refractive cornea (discussed next): these have a negative lens, enlarging the observed image by up to 50% over the receptor cells, thus increasing their optical resolution.[1]
[edit] Refractive cornea
Further information: Mammalian eye

In the eyes of most terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air, relieving the lens of the function of reducing the focal length. This has freed it up for fine adjustments of focus, allowing a very high resolution to be obtained.[1] As with spherical lenses, the problem of spherical aberration caused by the lens can be countered either by using an inhomogeneous lens material, or by flattening the lens.[1] Flattening the lens has a disadvantage: the quality of vision is diminished away from the main line of focus, meaning that animals requiring all-round vision are detrimented. Such animals often display an inhomogeneous lens instead.[1]

As mentioned above, a refractive cornea is only useful out of water; in water, there is no difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures which have returned to the water — penguins and seals, for example — lose their refractive cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strong cornea.[1]
[edit] Reflector eyes

An alternative to a lens is to line the inside of the eye with " mirrors", and reflect the image to focus at a central point.[1] The nature of these eyes means that if one were to peer into the pupil of an eye, one would see the same image that the organism would see, reflected back out.[1]

Many small organisms such as rotifers, copeopods and platyhelminths use such organs, but these are too small to produce usable images.[1] Some larger organisms, such as scallops, also use reflector eyes. The scallop Pecten has up to 100 millimeter-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses.[1]

There is at least one vertebrate, the spookfish, whose eyes include reflective optics for focusing of light. Each of the two eyes of a spookfish collects light from both above and below; the light coming from the above is focused by a lens, while that coming from below, by a curved mirror composed of many layers of small reflective plates made of guanine crystals.[19]
[edit] Compound eyes
Portrait of a Housefly (Musca domestica)
An image of a house fly compound eye surface by using Scanning Electron Microscope at X450 magnification
Arthropods such as this carpenter bee have compound eyes

A compound eye may consist of thousands of individual photoreception units. The image perceived is a combination of inputs from the numerous ommatidia (individual "eye units"), which are located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess a very large view angle, and can detect fast movement and, in some cases, the polarization of light.[20] Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained. This can only be countered by increasing lens size and number — to see with a resolution comparable to our simple eyes, humans would require compound eyes which would each reach the size of their head.

Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image.[21] Compound eyes are common in arthropods, and are also present in annelids and some bivalved molluscs.[22]

Compound eyes, in arthropods at least, grow at their margins by the addition of new ommatidia.[23]
[edit] Apposition eyes

Apposition eyes are the most common form of eye, and are presumably the ancestral form of compound eye. They are found in all arthropod groups, although they may have evolved more than once within this phylum.[1] Some annelids and bivalves also have apposition eyes. They are also possessed by Limulus, the horseshoe crab, and there are suggestions that other chelicerates developed their simple eyes by reduction from a compound starting point.[1] (Some caterpillars appear to have evolved compound eyes from simple eyes in the opposite fashion.)

Apposition eyes work by gathering a number of images, one from each eye, and combining them in the brain, with each eye typically contributing a single point of information.

The typical apposition eye has a lens focusing light from one direction on the rhabdom, while light from other directions is absorbed by the dark wall of the ommatidium. In the other kind of apposition eye, found in the Strepsiptera, lenses are not fused to one another, and each forms an entire image; these images are combined in the brain. This is called the schizochroal compound eye or the neural superposition eye. Because images are combined additively, this arrangement allows vision under lower light levels.[1]

Structure of the ommatidia of appositon compound eyes.
[edit] Superposition eyes

The second type is named the superposition eye. The superposition eye is divided into three types; the refracting, the reflecting and the parabolic superposition eye. The refracting superposition eye has a gap between the lens and the rhabdom, and no side wall. Each lens takes light at an angle to its axis and reflects it to the same angle on the other side. The result is an image at half the radius of the eye, which is where the tips of the rhabdoms are. This kind is used mostly by nocturnal insects. In the parabolic superposition compound eye type, seen in arthropods such as mayflies, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied decapod crustaceans such as shrimp, prawns, crayfish and lobsters are alone in having reflecting superposition eyes, which also has a transparent gap but uses corner mirrors instead of lenses.
[edit] Parabolic superposition

This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes.[7]
[edit] Other
The compound eye of a dragonfly

Good fliers like flies or honey bees, or prey-catching insects like praying mantis or dragonflies, have specialized zones of ommatidia organized into a fovea area which gives acute vision. In the acute zone the eyes are flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution.

There are some exceptions from the types mentioned above. Some insects have a so-called single lens compound eye, a transitional type which is something between a superposition type of the multi-lens compound eye and the single lens eye found in animals with simple eyes. Then there is the mysid shrimp Dioptromysis paucispinosa. The shrimp has an eye of the refracting superposition type, in the rear behind this in each eye there is a single large facet that is three times in diameter the others in the eye and behind this is an enlarged crystalline cone. This projects an upright image on a specialized retina. The resulting eye is a mixture of a simple eye within a compound eye.

Another version is the pseudofaceted eye, as seen in Scutigera. This type of eye consists of a cluster of numerous ocelli on each side of the head, organized in a way that resembles a true compound eye.

The body of Ophiocoma wendtii, a type of brittle star, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many chitons.
[edit] Relationship to lifestyle

Eyes are generally adapted to the environment and lifestyle of the organism which bears them. For instance, the distribution of photoreceptors tends to match the area in which the highest acuity is required, with horizon-scanning organisms, such as those that live on the African plains, having a horizontal line of high-density ganglia, while tree-dwelling creatures which require good all-round vision tend to have a symmetrical distribution of ganglia, with acuity decreasing outwards from the centre.

Of course, for most eye types, it is impossible to diverge from a spherical form, so only the density of optical receptors can be altered. In organisms with compound eyes, it is the number of ommatidia rather than ganglia that reflects the region of highest data acquisition.[1]:23-4 Optical superposition eyes are constrained to a spherical shape, but other forms of compound eyes may deform to a shape where more ommatidia are aligned to, say, the horizon, without altering the size or density of individual ommatidia.[24] Eyes of horizon-scanning organisms have stalks so they can be easily aligned to the horizon when this is inclined, for example if the animal is on a slope.[25] An extension of this concept is that the eyes of predators typically have a zone of very acute vision at their centre, to assist in the identification of prey.[24] In deep water organisms, it may not be the centre of the eye that is enlarged. The hyperiid amphipods are deep water animals that feed on organisms above them. Their eyes are almost divided into two, with the upper region thought to be involved in detecting the silhouettes of potential prey — or predators — against the faint light of the sky above. Accordingly, deeper water hyperiids, where the light against which the silhouettes must be compared is dimmer, have larger "upper-eyes", and may lose the lower portion of their eyes altogether.[24] Depth perception can be enhanced by having eyes which are enlarged in one direction; distorting the eye slightly allows the distance to the object to be estimated with a high degree of accuracy.[7]

Acuity is higher among male organisms that mate in mid-air, as they need to be able to spot and assess potential mates against a very large backdrop.[24] On the other hand, the eyes of organisms which operate in low light levels, such as around dawn and dusk or in deep water, tend to be larger to increase the amount of light that can be captured.[24]

It is not only the shape of the eye that may be affected by lifestyle. Eyes can be the most visible parts of organisms, and this can act as a pressure on organisms to have more transparent eyes at the cost of function.[24]

Eyes may be mounted on stalks to provide better all-round vision, by lifting them above an organism's carapace; this also allows them to track predators or prey without moving the head.[7]
[edit] Acuity
A hawk's eye

Visual acuity is often measured in cycles per degree (CPD), which measures an angular resolution, or how much an eye can differentiate one object from another in terms of visual angles. Resolution in CPD can be measured by bar charts of different numbers of white/black stripe cycles. For example, if each pattern is 1.75 cm wide and is placed at 1 m distance from the eye, it will subtend an angle of 1 degree, so the number of white/black bar pairs on the pattern will be a measure of the cycles per degree of that pattern. The highest such number that the eye can resolve as stripes, or distinguish from a gray block, is then the measurement of visual acuity of the eye.

For a human eye with excellent acuity, the maximum theoretical resolution is 50 CPD[26] (1.2 arcminute per line pair, or a 0.35 mm line pair, at 1 m). A rat can resolve only about 1 to 2 CPD.[27] A horse has higher acuity through most of the visual field of its eyes than a human has, but does not match the high acuity of the human eye's central fovea region.

Spherical aberration limits the resolution of a 7 mm pupil to about 3 arcminutes per line pair. At a pupil diameter of 3 mm, the spherical aberration is greatly reduced, resulting in an improved resolution of approximately 1.7 arcminutes per line pair.[28] A resolution of 2 arcminutes per line pair, equivalent to a 1 arcminute gap in an optotype, corresponds to 20/20 (normal vision) in humans.
[edit] Color

All organisms are restricted to a small range of the electromagnetic spectrum; this varies from creature to creature, but is mainly between 400 and 700 nm[29]. This is a rather small section of the electromagnetic spectrum, probably reflecting the submarine evolution of the organ: water blocks out all but two small windows of the EM spectrum, and there has been no evolutionary pressure among land animals to broaden this range.[30]

The most sensitive pigment, rhodopsin, has a peak response at 500 nm.[31] Small changes to the genes coding for this protein can tweak the peak response by a few nm;[2] pigments in the lens can also "filter" incoming light, changing the peak response.[2] Many organisms are unable to discriminate between colors, seeing instead in shades of "grey"; colour vision necessitates a range of pigment cells which are primarily sensitive to smaller ranges of the spectrum. In primates, geckos, and other organisms, these take the form of cone cells, from which the more sensitive rod cells evolved.[31] Even if organisms are physically capable of discriminating different colours, this does not necessarily mean that they can perceive the different colours; only with behavioral tests can this be deduced.[2]

Most organisms with colour vision are able to detect ultraviolet light. This high energy light can be damaging to receptor cells. With a few exceptions (snakes, placental mammals), most organisms avoid these effects by having absorbent oil droplets around their cone cells. The alternative, developed by organisms that had lost these oil droplets in the course of evolution, is to make the lens impervious to UV light — this precludes the possibility of any UV light being detected, as it does not even reach the retina.[31]:309
[edit] Rods and cones

The retina contains two major types of light-sensitive photoreceptor cells used for vision: the rods and the cones.

Rods cannot distinguish colors, but are responsible for low-light (scotopic) monochrome (black-and-white) vision; they work well in dim light as they contain a pigment, rhodopsin (visual purple), which is sensitive at low light intensity, but saturates at higher (photopic) intensities. Rods are distributed throughout the retina but there are none at the fovea and none at the blind spot. Rod density is greater in the peripheral retina than in the central retina.

Cones are responsible for color vision. They require brighter light to function than rods require. There are three types of cones, maximally sensitive to long-wavelength, medium-wavelength, and short-wavelength light (often referred to as red, green, and blue, respectively, though the sensitivity peaks are not actually at these colors). The color seen is the combined effect of stimuli to, and responses from, these three types of cone cells. Cones are mostly concentrated in and near the fovea. Only a few are present at the sides of the retina. Objects are seen most sharply in focus when their images fall on this spot, as when one looks at an object directly. Cone cells and rods are connected through intermediate cells in the retina to nerve fibers of the optic nerve. When rods and cones are stimulated by light, the nerves send off impulses through these fibers to the brain.[31]
[edit] Pigment

The pigment molecules used in the eye are various, but can be used to define the evolutionary distance between different groups, and can also be an aid in determining which are closely related – although problems of convergence do exist.[31]

Opsins are the pigments involved in photoreception. Other pigments, such as melanin, are used to shield the photoreceptor cells from light leaking in from the sides. The opsin protein group evolved long before the last common ancestor of animals, and has continued to diversify since.[2]

There are two types of opsin involved in vision; c-opsins, which are associated with ciliary-type photoreceptor cells, and r-opsins, associated with rhabdomeric photoreceptor cells.[32] The eyes of vertebrates usually contain cilliary cells with c-opsins, and (bilaterian) invertebrates have rhabdomeric cells in the eye with r-opsins. However, some ganglion cells of vertebrates express r-opsins, suggesting that their ancestors used this pigment in vision, and that remnants survive in the eyes.[32] Likewise, c-opsins have been found to be expressed in the brain of some invertebrates. They may have been expressed in ciliary cells of larval eyes, which were subsequently resorbed into the brain on metamorphosis to the adult form.[32] C-opsins are also found in some derived bilaterian-invertebrate eyes, such as the pallial eyes of the bivalve molluscs; however, the lateral eyes (which were presumably the ancestral type for this group, if eyes evolved once there) always use r-opsins.[32] Cnidaria, which are an outgroup to the taxa mentioned above, express c-opsins - but r-opsins are yet to be found in this group.[32] Incidentally, the melanin produced in the cnidaria is produced in the same fashion as that in vertebrates, suggesting the common descent of this pigment.[32]
[edit] See also
Wikipedia Books Wikipedia:Books has a book on: Eye

* Arthropod eye
* Human eye
* Mammalian eye
* Naked eye
* Sanpaku eyes

[edit] References

1. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af Land, M F; Fernald, R D (1992). "The Evolution of Eyes". Annual Review of Neuroscience 15: 1–29. doi:10.1146/annurev.ne.15.030192.000245. PMID 1575438.
2. ^ a b c d e Frentiu, Francesca D.; Adriana D. Briscoe (2008). "A butterfly eye's view of birds". BioEssays 30 (11-12): 1151. doi:10.1002/bies.20828. PMID 18937365.
3. ^ Nairne, James (2005). Psychology. Belmont: Wadsworth Publishing. ISBN 049503150x. OCLC 61361417. http://books.google.com/books?id=6MqkLT ... uNMCCFDdHw.
4. ^ Vicki Bruce, Patrick R. Green, and Mark A. Georgeson (1996). Visual Perception: Physiology, Psychology and Ecology. Psychology Press. pp. 20. ISBN 0863774504. http://books.google.com/books?id=ukvei0 ... o24zbrsfio.
5. ^ BioMedia Associates Educational Biology Site: What animal has a more sophisticated eye, Octopus or Insect?
6. ^ Who You Callin' "Shrimp"? – National Wildlife Magazine
7. ^ a b c d Cronin, T. W.; Porter, M. L. (2008). "Exceptional Variation on a Common Theme: the Evolution of Crustacean Compound Eyes". Evolution Education and Outreach 1: 463–475. doi:10.1007/s12052-008-0085-0. edit
8. ^ Halder, G., Callaerts, P. and Gehring, W.J. (1995). "New perspectives on eye evolution." Curr. Opin. Genet. Dev. 5 (pp. 602 –609).
9. ^ Halder, G., Callaerts, P. and Gehring, W.J. (1995). "Induction of ectopic eyes by targeted expression of the eyeless gene in Drosophila". Science 267 (pp. 1788–1792).
10. ^ Tomarev, S.I., Callaerts, P., Kos, L., Zinovieva, R., Halder, G., Gehring, W., and Piatigorsky, J. (1997). "Squid Pax-6 and eye development." Proc. Natl. Acad. Sci. USA, 94 (pp. 2421–2426).
11. ^ Conway-Morris, S. (1998). The Crucible of Creation. Oxford: Oxford University Press.
12. ^ Land, M.F. and Fernald, Russell D. (1992). "The evolution of eyes." Annu Rev Neurosci 15 (pp. 1–29).
13. ^ Eye-Evolution?
14. ^ a b Fernald, Russell D. (2001). The Evolution of Eyes: Where Do Lenses Come From? Karger Gazette 64: "The Eye in Focus".
15. ^ Kozmik, Zbynek; Ruzickova, Jana; Jonasova, Kristyna; Matsumoto, Yoshifumi; Vopalensky, Pavel; Kozmikova, Iryna; Strnad, Hynek; Kawamura, Shoji et al. (2008). "Assembly of the cnidarian camera-type eye from vertebrate-like components" (PDF). Proceedings of the National Academy of Sciences 105 (26): 8989–8993. doi:10.1073/pnas.0800388105. PMID 18577593. PMC 2449352. http://www.pnas.org/cgi/reprint/0800388105v1.pdf.
16. ^ Fernald, Russell D. (September 2006). "Casting a Genetic Light on the Evolution of Eyes". Science 313 (5795): 1914–1918. doi:10.1126/science.1127889. PMID 17008522.
17. ^ "Vision Optics and Evolution". BioScience 39 (5): 298–307. 1 May 1989. doi:10.2307/1311112. ISSN 00063568. edit
18. ^ a b c Wilson, M. (1978). "The functional organisation of locust ocelli". Journal of Comparative Physiology (4): 297–316.
19. ^ Wagner, H.J., Douglas, R.H., Frank, T.M., Roberts, N.W., and Partridge, J.C. (Jan. 27, 2009). "A Novel Vertebrate Eye Using Both Refractive and Reflective Optics". Current Biology 19 (2): 108–114. doi:10.1016/j.cub.2008.11.061. PMID 19110427.
20. ^ Völkel, R; Eisner, M; Weible, K. J (June 2003). "Miniaturized imaging systems" (PDF). Microelectronic Engineering 67-68 (1): 461–472. doi:10.1016/S0167-9317(03)00102-3. http://www.suss-microoptics.com/downloa ... ystems.pdf.
21. ^ Gaten, Edward (1998). "Optics and phylogeny: is there an insight? The evolution of superposition eyes in the Decapoda (Crustacea)". Contributions to Zoology 67 (4): 223–236. http://dpc.uba.uva.nl/ctz/vol67/nr04/art01#FIGURE1.
22. ^ Ritchie, Alexander (1985). "Ainiktozoon loganense Scourfield, a protochordate? from the Silurian of Scotland". Alcheringa 9: 137. doi:10.1080/03115518508618961.
23. ^ Mayer, G. (2006). "Structure and development of onychophoran eyes: What is the ancestral visual organ in arthropods?". Arthropod Structure and Development 35 (4): 231–245. doi:10.1016/j.asd.2006.06.003. PMID 18089073.
24. ^ a b
I like the Arcitc Monkeys....no really, I do!
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Postby noonsun » Sun Dec 20, 2009 2:52 am

Well done, Sherlock
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Postby Eddie » Sun Dec 20, 2009 3:04 am

Elementary, my dear Watson.
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